Introduction. Populus trichocarpa TORR. and GRAY. By M. G. R. CANNELL and S. C. WILLETT

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1 Shoot Growth Phenology, Dry Matter Distribution and Root: Shoot Ratios of Provenances of Populus trichocarpa, Picea sitchensis and Pinus contorta growing in Scotland By M. G. R. CANNELL and S. C. WILLETT Institute of Terrestrial Ecology, Bush Estate, Penicuik, Midlothian EH26 WB, Scotland (Received April / July 1976) Introduction Inherent differences in root : shoot weight ratios of trees are of interest as factors that may influence wind stability, and because they may reflect differences in the relative growth rates of the roots and shoots. Windthrow, basal stem bending and wind-loosening are serious problems in Britain (EDWARDS et al., 1963; LINES and BOOTH, 1972) and any genetic gain in wind stability would be valuable. It is assumed in this paper that genotypes which have relatively heavy root systems (and hence proportionately smaller shoots and sail area) will often be better anchored than those with light root systems, although there will, of Course, be differences in root morphology and responses to soil conditions. In the case of Pinus contorta there is good evidence that southerly, fast-growing provenances, which are often not wind-firm in Britain, have particularly small root : shoot weight ratios both as nursery seedlings and 8-year-old forest trees (LINES, 1971 and pers. comm.). The relative growth rates of roots and shoots are important parameters of dry matter producion. They reflect, for instance, the proportion of total dry matter gain that is reinvested in photosynthetic tissue. The functional, hormone-mediated equilibrium that governs the partition of assimilates between shoots and roots is at the core of most models of plant growth (e.g. BROUWER, 1962; LEDIG, 1969; DE WIT et al., 1971; PROMNITZ, 1975). The ratio of the relative growth rates of shoots and roots in young plants with exponential growth is given approximately by the 'allometric coefficient' k in the equation: log shoot dry weight = a + k log root dry weight. If k is greater than 1.0 the root : shoot ratio will decrease with increase in plant weight, and if it is less than 1.0 the root : shoot ratio will increase with increase in plant weight (LEDIG et al., 1970; WAREING and PATRICK, 1975). The value of k is governed by the relative 'activity' of the root and shoot masses, such that root weight X rate (absorption) leaf weight X rate (photosynthesis). Most herbaceous plants will respond to a change in root or shoot 'activity' by a compensating change in root and shoot masses (THORNLEY, 1972; HUNT, 1975; also THORNLEY, 1975). MAGGS (1961) working with young apple rootstocks and LEDIG et al. (1970), reviewing work on conifer seedlings, agreed that k could differ between tree genotypes, but stressed that k was unusually stable in trees and only altered by drastic environmental treatments or changes in physiology. Such physiological changes undoubtedly include the temperature and photoperiodically induced changes in the activity of the shoot apices, since k can only remain constant as long as both the shoots and roots are competing dry matter sinks. Thus, WAREINC (1950) showed that k decreased in Pinus sylvestris seedlings that had stopped elongating in short days for several weeks; that is, their roots then grew relatively faster than their shoots. The Same phenomenon is evident in HEIDE'S (1974) data on dry weight changes in Picea abies seedlings in different photoperiodic regimes. Also, SWEET and WAREING (1968) working on Pinus contorta seedlings, and SCHULTZ and GATHERUM (1971) on Pinus sylvestris, showed that provenances which were photoperiodically induced to stop elongating their shoots early in their first growing season subsequently developed relatively large root : shoot dry weight ratios compared with provenances which continued elongating late into the autumn. This paper describes observations extending those already made by SWEET and WAREING, and SCHULTZ and GATHERUM. DO provenances of contrasting species with similar natural ranges show similar geographic trends in root : shoot ratios? Are provenance differences in root : shoot ratio linked with differences in seasonal periods of shoot growth? And do differences in shoot growth phenology alter the values of k as suggested by WAREING (1950) and HEIDE (1974), and, if so, are the resulting differences in root : shoot ratio compounded over successive seasons? The three species analysed, Populus trichocarpa, Picea sitchensis and Pinus contorta, all have natural ranges along the western seaboard of North America and are shown to have comparable photoperiodic ecotypes. They are all grown commercially in Britain, where some provenances are clearly ill-adapted to the mild, maritime climate with northern daylength mgimes and strong winds. Populus trichocarpa TORR. and GRAY Periods of shoot elongation Populus trichocarpa ranges from Alaska to California and inland to the Rocky Mountains and Montana. The dates when clones from throughout this range stopped growing in height between June and October 1951 at Weston, Mass., U.S.A. (42O 22' N) were inversely correlated with their latitudes of origin (r = -0.82) and the dates for clones from latitudes 45 to 47O N (where climatic data were available) were positively correlated with the lengths of the growing seasons at their places of origin ( days, r = 0.79; PAULEY and PERRY, 1954). A collection of 98 clones of Populus trichocarpa from 62 locations throughout its range were established in unreplicated rows in a nursery at the Bush Estate, Midlothian, Scotland, in 1972 (55O 50' N). Some cuttings had been taken from PAULEY'S surviving collection at Weston with generous help from the Maria Moors Cabot Foundation, and others had been kindly sent by workers in western North Ameri- Ca. By taking weekly height measurements on 5 plants during 1973, the date was estimated when each clone completed 95% of its total height growth. PAULEY and PERRY'S findings were confinned, and using Smithsonian tables of global Silvae Genetica 25, 2 (1976)

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